Patterns are orders embedded in randomness. They may appear as spatial arrangements or temporal series, and the elements may appear identical or with variations. Patterns exist in the physical world as well as in living systems. In the biological world, patterns can range from simple to complex, forming the basic building blocks of life. The process which generates this ordering in the biological world was termed pattern formation. Since Wolpert promoted this concept four decades ago, scientists from molecular biology, developmental biology, stem cell biology, tissue engineering, theoretical modeling and other disciplines have made remarkable progress towards understanding its mechanisms. It is time to review and re-integrate our understanding. Here, we explore the origin of pattern formation, how the genetic code is translated into biological form, and how complex phenotypes are selected over evolutionary time. We present four topics: Principles, Evolution, Development, and Stem Cells and Regeneration. We have interviewed several leaders in the field to gain insight into how their research and the field of pattern formation have shaped each other. We have learned that both molecular process and physico-chemical principles are important for biological pattern formation. New understanding will emerge through integration of the analytical approach of molecular-genetic manipulation and the systemic approach of model simulation. We regret that we could not include every major investigator in the field, but hope that this Special Issue of the Int. J. Dev. Biol. represents a sample of our knowledge of pattern formation today, which will help to stimulate more research on this fundamental process.
KEY WORDS: development, evolution, Evo-Devo, systems biology, morphogenesis, pattern formationWhen we see patterns in peacock feathers (Fig. 1), leopard spots (Murray, 2003;Liu et al., 2006), or zebra stripes, we are fascinated by the order, the variations and the beauty. The biological processes that generate this ordering are pattern formation, a term popularized by Wolpert to describe the spatial organization of cell differentiation (Wolpert, 1969, and interview of Wolpert in Richardson, 2009a). Indeed, the processes of pattern formation can occur in the physical world as well as in living systems (Kauffman, 1993;Ball, 1999). Thus, we can also appreciate patterns in water waves, rock layers, and sand dunes (Fig. 2, Hazen, 2009), which emerge from apparent chaos or randomness. In living organisms, we can see patterns at different hierarchical levels, from molecular pathways, genome organization, developmental interactions, tissues, organs, body plans, and also at the level of animal behaviors. While the possibility of disorganization and disintegration is always present, organized patterns are able to emerge, and become stable, in many different systems and contexts. The formation of patterns is one of the key properInt. J. Dev. Biol. 53: 653-658 (2009)
ties of life.What is the origin of the pattern-organizin...